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A Lagrangian Study of Precipitation-Driven Downdrafts*
GIUSEPPE TORRI
Department of Earth and Planetary Sciences, Harvard University, Cambridge, Massachusetts
ZHIMING KUANG
Department of Earth and Planetary Sciences, and School of Engineering and Applied Sciences,
Harvard University, Cambridge, Massachusetts
(Manuscript received 29 July 2015, in final form 9 November 2015)
ABSTRACT
Precipitation-driven downdrafts are an important component of deep convective systems. They stabilize
the atmosphere by injecting relatively cold and dry air into the boundary layer. They have also been invoked
as responsible for balancing surface latent and sensible heat fluxes in the heat and moisture budget of tropical
boundary layers. This study is focused on precipitation-driven downdrafts and basic aspects of their dynamics
in a case of radiative–convective equilibrium. Using Lagrangian particle tracking, it is shown that such
downdrafts have very low initial heights, with most parcels originating within 1.5 km from the surface. The
tracking is also used to compute the contribution of downdrafts to the flux of moist static energy at the top of
the boundary layer, and it is found that this is on the same order of magnitude as the contribution due to
convective updrafts, but much smaller than that due to turbulent mixing across the boundary layer top in the
environment. Furthermore, considering the mechanisms driving the downdrafts, it is shown that the work
done by rain evaporation is less than half that done by condensate loading.
1. Introduction
The existence of downdrafts in deep convective sys-
tems was acknowledged as early as a hundred years ago
by Humphreys (1914), who also hypothesized that they
are maintained by the evaporation of rain falling
through unsaturated air. Since then, downdrafts have
been the focus of many studies (e.g., Knupp and Cotton
1985; Cotton et al. 2011, and references therein), which
have led to a considerable advancement of our un-
derstanding of these phenomena and, more generally, of
precipitating convection.
Within the category of downdrafts, we would like to
concentrate on precipitation-driven downdrafts, bywhich,
following the nomenclature explained in Knupp and
Cotton (1985), wemean those downdrafts that appear in
the presence of precipitation and that reach the surface,
injecting masses of cold and dry air in the subcloud layer
(e.g., Byers and Braham 1949; Fankhauser 1976; Barnes
and Garstang 1982; Knupp 1987, 1988).
From a rather practical point of view, precipitation-
driven downdrafts have been of interest for some time,
given their potential to generate downbursts and mi-
crobursts, very strong currents of air with vertical ve-
locities often exceeding 10m s21 in magnitude (see, e.g.,
Fujita 1985, 1986; Proctor 1988, 1989; Wakimoto et al.
1994). These intense downdrafts can be found in cases
where precipitating clouds are formed above deep and
dry boundary layers—such as over the Great Plains—
and constitute a serious hazard for civil aviation.
From a more conceptual perspective, throughout the
years precipitation-driven downdrafts have been studied
in different contexts, an important example being that of
the so-called boundary layer quasi-equilibrium (BLQE)
hypothesis (Emanuel 1989; Raymond 1994, 1995), pro-
posed to explain why the moist entropy of tropical
boundary layers seems to change very little with time.
Simply put, according to this hypothesis, the moistening
and heating of the tropical boundary layer provided by
* Supplemental information related to this paper is available at
the Journals Online website: http://dx.doi.org/10.1175/JAS-D-15-
0222.s1.
Corresponding author address: Giuseppe Torri, Department of
Earth and Planetary Sciences, Harvard University, 20 Oxford
Street, Cambridge, MA 02138.
E-mail: [email protected]
FEBRUARY 2016 TORR I AND KUANG 839
DOI: 10.1175/JAS-D-15-0222.1
� 2016 American Meteorological Society
the surface fluxes are counterbalanced mainly by
precipitation-driven downdrafts that bring low-entropy
air from the midtroposphere to the surface. As argued by
Raymond (1995), the turbulent entrainment at the top of
the boundary layer plays little role in the balance. This
explanation is in contrast with the hypothesis used by
Arakawa and Schubert (1974) in their cumulus parame-
terization, where it is mixing with the environmental air
and, sometimes, horizontal advection, which balance the
action of surface fluxes. These two hypotheses were re-
cently examined and tested by Thayer-Calder and
Randall (2015) with a numerical model. Interestingly,
their results suggest that only a small fraction of the flux
of moist static energy (MSE) at the top of the boundary
layer is due to precipitation-driven downdrafts.
Another framework in which precipitation-driven
downdrafts are of considerable importance is that of
convection triggering. The masses of air injected by
downdrafts in the subcloud layer spread horizontally on
the surface as density currents, usually called cold pools,
which are one of the main factors responsible for
maintaining deep convective systems (Purdom 1976;
Weaver and Nelson 1982). In organized cases, such as
squall lines or supercell thunderstorms, it has been
known for a long time (see, e.g., Rotunno and Klemp
1985; Rotunno et al. 1988; Weisman et al. 1988;
Weisman and Rotunno 2004) that the interaction be-
tween the gust front of cold pools and the wind shear or
the collision between two or more cold pools can pro-
vide enough kinetic energy to parcels at the surface to
reach their level of free convection (LFC). In non-
organized cases, parcels reach their LFC through a co-
operation of the gust-front lifting and the thermodynamic
effect provided by the positive moisture anomaly at the
leading edge of cold pools, which reduces the convective
inhibition experienced by parcels above the lifting con-
densation level (Tompkins 2001; Torri et al. 2015). The
influence of precipitation-driven downdrafts on convec-
tion triggering is indirect but, nevertheless, strong: by
controlling the initial conditions of cold pools, downdrafts
effectively control the strength of the cold pools’ gust
fronts and their ability to lift parcels from the surface
and, potentially, also the magnitude of the thermo-
dynamic effect.
In this study, we will consider a tropical maritime
environment, and we will focus on three fundamental
aspects of precipitation-driven downdraft dynamics:
1) What is the initial height of a precipitation-driven
downdraft?
2) What is the contribution of precipitation-driven
downdrafts to the budget of MSE in the subcloud
layer?
3) What is the role of condensate loading versus rain
evaporation in maintaining a precipitation-driven
downdraft?
To some extent, these topics have been addressed in
the past in various studies, although using limited tech-
niques that did not provide an entirely clear answer. For
instance, starting from as early as Mal and Desai (1938),
the typical approach to diagnose the initial height of
downdrafts, sometimes referred to as source level, has
been to use thermodynamic properties of downdraft air,
like the wet-bulb potential temperature uw, the equiva-
lent potential temperature ue, or MSE, which are ap-
proximately conserved in adiabatic processes and water
vapor–liquid water phase transitions (see also, e.g., Mal
and Desai 1938; Normand 1946; Newton 1950; Zipser
1969; Betts 1976; Fankhauser 1976; Lemon 1976; Barnes
and Garstang 1982; Johnson and Nicholls 1983;
Kingsmill and Houze 1999). The problem with this ap-
proach is that it assumes that mixing is negligible and
that all the downdraft air originates at the initial height,
neither of which hypotheses are necessarily true.
Analogously, various authors have asked what
mechanisms maintain precipitation-driven downdrafts,
recognizing rain evaporation and condensate loading as
leading factors (Byers and Braham 1949; Braham 1952;
Fankhauser 1976; Lemon 1976; Barnes and Garstang
1982; Knupp and Cotton 1985). However, the extent to
which one mechanism is dominant over the other even in
simple cases of unorganized convection remains unclear.
In this manuscript, we will address the two questions
outlined above mainly using a Lagrangian particle dis-
persion model (LPDM). Compared to the techniques
that have been used in the past, the LPDM provides a
great deal of information. In particular, analyzing the
position of millions of particles released in the simula-
tion domain will allow us to determine the initial height
of a downdraft in a clear and simple manner. Also,
combining the output from the LES model that we will
use with the Lagrangian history of the particles from
when they enter a precipitation-driven downdraft until
their exit near the surface, we will be able to quantify the
importance of rain evaporation and condensate loading
in driving the downdraft.
Notice that the use of particles’ trajectories to study
downdrafts is a path that has been explored in the past:
Miller and Betts (1977), for example, used them to il-
lustrate the behavior of downdrafts in Venezuelan
convective storms, and Knupp (1988) used them in the
context of storms in the high plains. However, the
computational limitations of the past did not allow for
the simulation of a number of trajectories large enough
to make significant statistics.
840 JOURNAL OF THE ATMOSPHER IC SC IENCES VOLUME 73
2. Methods
The model used in this study is the System for At-
mospheric Modeling (SAM), version 6.8.2, which solves
the anelastic equations of motion and uses liquid water
static energy and nonprecipitating and precipitating
total water as thermodynamic prognostic variables
(Khairoutdinov and Randall 2003). The equations are
solved with doubly periodic boundary conditions in the
horizontal directions. A prognostic turbulent kinetic
energy 1.5-order closure scheme is used to parameterize
subgrid-scale effects.
We ran the model with the Lin microphysics scheme,
a single-moment scheme that predicts the specific hu-
midities of water vapor, cloud liquid water, cloud ice,
rain, snow, and graupel (Lin et al. 1983). The surface
fluxes are computed using the Monin–Obukhov simi-
larity theory.
The simulations are carried out over a domain of 64364km2 in the horizontal directions and 30 km in the
vertical. The grid size in the horizontal directions is
250m, and it varies in the vertical: 500m above 10 km,
which decrease progressively to 38m close to the model
surface. The time step is 3 s. Figure 1 shows the envi-
ronment vertical profiles of potential temperature
(red), water vapor specific humidity (blue), and relative
humidity (green), averaged over the duration of the
simulation.
We ran the model with the LPDM discussed in Nie
and Kuang (2012). For every column in the domain, the
LPDM is initialized with 1120 particles, their positions
being distributed randomly over the bottom 18kmof the
model domain, with a probability distribution that is a
uniform function of pressure. A validation of the LPDM
can be found in the supplemental information.
Every minute of simulated time, the positions of the
Lagrangian particles and the three-dimensional model fields
fromSAMare recorded.A cross-comparison of Lagrangian
and Eulerian data provides the complete histories of ther-
modynamic and dynamic properties of the particles.
Throughout the simulation, the sea surface tempera-
ture is held constant at 302.65K, and the diurnal cycle is
removed by fixing the zenith angle at 51.78 with solar
constant halved to 685Wm22. The mean winds are ini-
tialized to zero. We first ran the model for 30 days
without Lagrangian particles on the same domain until
radiative–convective equilibrium (RCE) is reached. At
the end of this period, we restarted the simulation with
the Lagrangian particles and collected data for 12 h of
model time. No nudging was imposed on the horizontal
winds. The precipitation rate averaged in time during
the 12h of data collection and in space over the entire
domain is 3.4mmday21.
We define the subcloud layer as the portion of the
model domain that includes the surface and in which
the time- and domain-averaged cloud liquid water
specific humidity ql is below 1025 g kg21. With the
chosen settings, this corresponds to the bottom 531m
of the domain.
We define the density potential temperature of a
particle as follows (Emanuel 1994):
ur5 u
�11
�R
y
Rd
2 1
�qy2 q
l
�, (1)
where Rd and Ry are the gas constants of dry air and
water vapor and equal 287 and 461 J kg21K21, re-
spectively, and qy is the specific humidity of water vapor.
As customary, we define «5 (Ry/Rd 2 1) and will use
this convention for the remainder of this manuscript.
We say that a grid box in the subcloud layer is part of
the core of a cold pool if its density potential
FIG. 1. Time average of the vertical profiles of potential tem-
perature (red), water vapor specific humidity (blue), and relative
humidity (green).
FEBRUARY 2016 TORR I AND KUANG 841
temperature ur is lower than the horizontal average by
1.5K.1 This definition was chosen after examining the
model outputs and only serves to separate the part of the
cold pool that is fed by the precipitation-driven down-
draft from the rest (cf. Torri et al. 2015). As shown in the
supplemental information, the analysis presented in the
following section is not particularly sensitive to the value
chosen for the threshold.
We consider as part of a precipitation-driven downdraft
all particles from any altitude that acquire and maintain
negative vertical velocity until they enter the core of a
cold pool. Interestingly, although we do not require the
precipitation-driven downdraft to be unsaturated, most of
the descent of the particles in precipitation-driven down-
drafts takes place in unsaturated grid boxes.
3. Results
Having laid out the research questions in section 1,
and after having explained how we pursue these ques-
tions in the section 2, we will now discuss our findings.
a. On the initial height of precipitation-drivendowndrafts
Following the definition of precipitation-driven down-
draft that we have given in the previous section,
whenever a particle enters the core of a cold pool, we
track its position back in time for every step during which
the particle was in grid boxes that satisfied the condition
that the vertical wind w# 0ms21. The height of the first
grid box for which the previous condition is true is taken
as the initial height of the particle we sampled.
Notice that, because the coldest regions of cold pools
are known to exhibit high pressure perturbations, the
model outputs show that the vertical velocity of
precipitation-driven downdrafts as they approach the
surface decreases considerably (cf. Fig. 10 later in the
text). When the vertical velocity is small enough, tur-
bulent or wave fluctuations may cause its values in some
grid boxes to oscillate between positive and negative.
There follows that, if we were to apply the definition of
precipitation-driven downdraft given above, a consid-
erable number of sampled particles would appear to
have originated close to the surface. To avoid this
problem, whenever the vertical velocity of a particle in a
downdraft turns positive, a clock is started as the parti-
cles’ positions are scanned backward in time; if the
vertical velocity becomes negative again within 10min
from the start of the clock, the particle is still considered
part of the downdraft; otherwise, it is dismissed. Sensi-
tivity tests presented in the supplemental information
show that the results and our conclusions are not par-
ticularly sensitive to the precise value chosen for the
time threshold. Furthermore, in order to avoid in-
terference from fast detrainment–entrainment events in
cold pools, once a particle ends up in a cold pool core
and its properties are recorded, the particle is flagged
and discarded for the rest of the simulation.
In Fig. 2, the blue curve shows the distribution of initial
heights of all particles that were injected in the core of a
cold pool by a precipitation-driven downdraft. Notice that
the distribution is confined almost entirely to the bottomof
the troposphere: 98%of particles have initial heights lower
than 2.5km. The curve shows a pronounced peak at ap-
proximately 1km of altitude, just a few hundred meters
above the top of the subcloud layer, and a smaller one near
the surface. Examination of trajectories for particles
that contribute to the lower peak shows that most of
these particles descended from higher altitudes with a
precipitation-driven downdraft, but, near the surface, their
vertical velocity, however small, was positive formore than
10min before they entered the core of a cold pool.
These results are nicely in accordance with existing
literature, which suggested that, for tropical convection,
the source level of precipitation-driven downdrafts is just
FIG. 2. Distribution of Lagrangian particles’ positions at various
times before becoming part of a precipitation-driven downdraft.
The red, green, and black curves refer to positions 30, 60, and
90min before entering the downdraft, respectively. The blue curve
shows the distribution of positions at the time of entrance.
1 For brevity, whenever a particle enters a grid box that is part of
the core of a cold pool, we will simply say that ‘‘the particle enters/
belongs to the core of a cold pool.’’
842 JOURNAL OF THE ATMOSPHER IC SC IENCES VOLUME 73
above cloud base [see, e.g., Zipser (1969), Betts (1976),
and Kingsmill and Houze (1999), just to cite a few].
We should point out that the initial height of a
downdraft may still carry along some ambiguity. For
example, it could be that a number of particles that
entered the precipitation-driven downdraft at 1 km ac-
tually originated frommuch greater altitudes: they were
transported down by another downdraft from which
they eventually left, and then, after some time, they
became part of the precipitation-driven downdraft that
carried them down to the surface. A simple way to check
whether this is the case is to ask where the particles were
before entering the precipitation-driven downdraft.
The red, green, and black curves in Fig. 2 show the
distribution of the sampled particles’ positions 30, 60,
and 90min before entering the precipitation-driven
downdraft, respectively. They can be thought of as the
evolution back in time of the blue curve. Interestingly,
the red curve suggests that 61% of particles actually
come from the subcloud layer, whereas the remaining
seem to originate near the height where they entered the
downdraft. According to the nomenclature discussed in
Knupp (1988), we will refer to the former as up–down
and to the latter asmidlevel. Less than 2%of all particles
originate from an altitude greater than 2.5 km.
To better appreciate the history of the Lagrangian par-
ticles prior to their entrance in a precipitation-driven
downdraft, we can look directly at the trajectories of a few
representative cases. The red lines in Fig. 3 show the time
evolution of the vertical positions of three particles for the
60min before they enter the core of a cold pool. Notice that
the horizontal axis represents the time in the particles’ his-
tories, with 0min being themoment they enter a cold pool’s
core. The particles have been selected rather arbitrarily, but
an extensive scan through a large number of particles’ his-
tories makes us confident the particular examples shown
illustrate well the possible scenarios for downdrafts.
In all panels, the blue contours represent the total
nonprecipitating water specific humidity qn, whereas the
green contours represent the buoyancy. To keep the image
clean, we only show contours for negative buoyancy and
only for the subcloud layer. Finally, the gray shading rep-
resents the total precipitating water specific humidity qp.
Figure 3 (left) shows the history of a particle in an up–
down downdraft. As can be seen, the particle is lifted from
the surface at approximately225min by the gust front of
an expanding cold pool, indicated by the appearance of
the negative buoyancy contours in the subcloud layer, and
it quickly becomes part of an updraft. As the particle as-
cends, it reaches lifting condensation level (LCL), and
soon afterward it enters a precipitating column. The pre-
cipitation experienced by the particle is sufficient to
change the sign of its buoyancy and stop its ascension.As a
result, the particle leaves the updraft and enters a down-
draft that becomes unsaturated almost immediately. The
descent continues with no interruptions until the particle
reaches the core of a cold pool. Curiously, visual in-
spection of the three-dimensional model output suggests
that the particle enters the same cold pool that had lifted it
from the surface.
FIG. 3. Three examples of Lagrangian particles’ histories 60min before entering the core of a cold pool are shown with a red line. (left)
An up–down downdraft, (middle) a midlevel downdraft originating at ;1 km, and (right) a hybrid between the two. The blue contours
represent the specific humidity of total nonprecipitating water at increasing intervals of 1 g kg21. The green contours show negative
buoyancy in the subcloud layer at decreasing intervals of 0.02m s22 starting at 20.01m s22. The gray shading represents the specific
humidity of total precipitating water. The values shown for all variables at different heights refer to the vertical column where the
particle is.
FEBRUARY 2016 TORR I AND KUANG 843
Figure 3 (middle) shows the history of a particle that
descends to the surface levels through a midlevel
downdraft. As might be expected, the particle sits in an
almost undisturbed environment for most of its life. The
changes in altitude in clear sky seem to be due to ran-
dom turbulent fluctuations or gravity waves. Then, at
approximately 220min, the particle is entrained into a
cloud, where its buoyancy becomes negative. The par-
ticle starts descending and enters the core of a cold pool
at the surface.
Finally, Fig. 3 (right) shows another possible scenario
for particles entering precipitation-driven downdrafts—
one that, to some extent, is a mixture of the previous two.
The particle represented in the figure starts off at an al-
titude of approximately 1km, subject only to random
turbulence or wave motions, which cause its position to
oscillate a little. At220min, the particle becomes part of
an updraft that makes it rise to an altitude of 2km, where,
finally, its buoyancy reverses sign and the particle enters
an unsaturated downdraft all theway down to the surface.
This strengthens the conclusion drawn from Fig. 2 that
particles with higher initial height may actually originate
atmuch lower altitude and have, therefore, a higherMSE
than what the initial height could suggest.
One of the messages that emerges from the previous
discussion is that precipitation-driven downdrafts tend to
have relatively low initial heights, possibly transporting
parcels that originated in the bottom kilometer of the
troposphere. To provide further evidence to support this
statement, we will use three methods, one based on the
Lagrangian and two on the Eulerian point of view.
The former one consists of changing the sampling
technique we have adopted above, and, instead of con-
sidering particles that enter a downdraft and then reach
the surface layers inside the core of a cold pool, we
simply track all particles in downdrafts, and we follow
them until they leave it, regardless of the height where
this happens. To be more specific, whenever a particle
enters a grid box in which w # 20.5m s21, its positions
are saved for as long as the particle is in a grid box with
negative vertical velocity (i.e., w # 0ms21).
For every particle, we collect the initial height and the
final height, which, in contrast to the previous sampling
of precipitation-driven downdrafts, does not necessarily
have to be in a cold pool or even in the subcloud layer.
We also do not allow for a temporal window between
the time a particle leaves the downdraft and when its
final height is recorded, as done with the other sam-
pling technique. Nevertheless, in the supplemental in-
formation we show that considering such a window has
no effect on our conclusions.
Next, we define the height matrix as the matrix where
the (i, j) element contains the density of particles per
meter square that entered a downdraft at height zj and
left it at height zi. Figure 4 shows the logarithm of this
matrix. As a guiding rule, numbers in the same column
refer to particles that became part of a downdraft at the
same height.
The figure suggests that, although there are down-
drafts at all heights, they do not travel much farther
than a few hundred meters. The reason for this, as also
suggested by others (see, e.g., Böing et al. 2014), is that,
as downdraft parcels descend over large distances, they
become dry, and, thus, they start following a dry adiabat.
Because the environment above the subcloud layer re-
mains close to a moist adiabat, these parcels quickly lose
their negative buoyancy. Furthermore, because pre-
cipitating columns are not steady, the negative contri-
butions of rain evaporation and condensate loading to
the parcels’ buoyancy are also going to become in-
sufficient to maintain the parcels’ descent.
One of the most curious features of Fig. 4 that jumps
out at the viewer is that downdrafts originating between
3 and 5km seem to be able to travel a greater distance
than those originating at other heights. The average
profile of rain evaporation rate shown by the blue curve
in Fig. 5, and that of the graupel melting rate shown in
green, suggests that particles in downdrafts that cross
the levels between 2 and 4km would experience signif-
icant evaporation of rain and melting of graupel, which
could provide extra kinetic energy to travel a longer
FIG. 4. Logarithm of the height matrix, the (i, j) entry of which
indicates the number of particles that became part of a downdraft
at height zj and exited at height zi.
844 JOURNAL OF THE ATMOSPHER IC SC IENCES VOLUME 73
distance than particles that descended frommuch higher
or lower altitudes. Notice that, although the peak in
melting rate of graupel is almost an order of magnitude
higher than the rain evaporation rate at the same alti-
tude, the latent heat of fusion is roughly a factor of 8
smaller than the latent heat of vaporization, so melting
of graupel is less efficient in cooling the parcels, and thus
in driving the downdraft, than rain evaporation. Finally,
we should note that the minimum of evaporation rate
between 1 and 2km is the result of high relative hu-
midity, and thus low saturation deficit, at these altitudes.
The second consistency check that we perform is
based on the distribution of MSE in cold pool cores, by
which, we remind the reader, we mean those grid cells
with a ur anomaly smaller than 21.5K. Because adia-
batic processes and phase transitions leaveMSE roughly
unchanged, and because MSE is monotonically de-
creasing in approximately the bottom 5km of the tro-
posphere, the presence of low-MSE grid boxes in cold
pool cores would be the signature of air parcels origi-
nating in the midtroposphere. The red curve in Fig. 6
shows the cumulative distribution function of MSE in
cold pool cores during the entire simulation, whereas the
blue curve represents the temporal and spatial average
of the environmental profile of MSE in the bottom 5km
of the troposphere. The cumulative distribution is con-
centrated around relatively large values of MSE: the 1st
percentile corresponds to an MSE of 333.8K, which
Fig. 6 indicates as the average value in the environment
at an altitude of 1.3 km. Notice that MSE is not con-
served under mixing, so our estimates are likely to be
biased low. Nevertheless, such low altitudes as those
indicated above support the claim that air in cold pool
cores is unlikely to originate in the midtroposphere.
Finally, the third method that we use to check the
consistency of our claim that initial heights of precipitation-
driven downdrafts are very low is based on the joint
distribution of MSE and vertical velocity at a given
height. In particular, Fig. 7 (left) shows the logarithm
of the time-averaged joint distribution of MSE
anomaly—computed at each time step with respect to
the horizontal average—and the vertical velocity at
531m near the top of the subcloud layer. The black
curve highlights the contour corresponding to the
value 4.5 of the logarithm of the distribution, which
encloses 99.7% of the total number of grid boxes
sampled and 87.5% of the total flux of MSE. The
contour line shows that most of the MSE flux is given
by grid boxes with an MSE anomaly higher than 26K
and relatively small vertical velocities, which suggests
that downdrafts from the midtroposphere do not inject
much air in the boundary layer.
FIG. 5. Time averages of the vertical profiles of rain evaporation
rate (blue), graupelmelting rate (green), and total precipitating water
specific humidity (red) for the bottom 5km of the model domain.
FIG. 6. Time average of the vertical profile of MSE in the envi-
ronment (blue) and cumulative distribution function of MSE in
cold pool cores (red).
FEBRUARY 2016 TORR I AND KUANG 845
b. On the contribution of precipitation-drivendowndrafts to the subcloud-layer MSE
At this point, it is natural to try and quantify the role
of the various transport categories—updrafts, down-
drafts, and the environment—in balancing the action of
surface fluxes in the subcloud layer. To begin with, let us
consider a column of air of unit surface and define the
mass contained in the subcloud-layer portion of the
column as
[r]5
ðztopzsfc
r dz , (2)
where ztop and zsfc are the heights of the top of the
subcloud layer and the surface. Because mass can be
exchanged between the subcloud layer and the free
troposphere, if we call Mi the mass flux associated with
the i transport category, the temporal variation in mass
per unit surface in the subcloud layer can then be written
as follows:
›[r]
›t5 �
i2TM
i, (3)
where T5 fupdraft, downdraft, environmentg. Along
the same lines of reasoning, the interaction of the sub-
cloud layer and the free troposphere causes the total
MSE per unit surface in the subcloud layer, defined by
[rh]5
ðztopzsfc
(rh) dz , (4)
to vary in time according to the following:
›[rh]
›t5 �
i2TM
ihi, (5)
where hi is the value of MSE at the top of the subcloud
layer associated with the transport category i. Notice
that the subcloud-layer MSE can also change because of
radiative effects, which we will represent as QR, and
through sensible and latent heat fluxes at the surface,
which we will write as SH and LH. It follows that the
complete form of Eq. (5) is given by
›[rh]
›t5 �
i2TM
ihi1 (SH1LH)1 [rQ
R] , (6)
where [rQR] is defined in a similar fashion to [rh]. If we
now assume that the air in the subcloud layer obeys the
Boussinesq approximation and has constant density r0,
we can rewrite the left-hand side of the above equation
as
›[rh]
›t5
ðztopzsfc
›r
›th1 r
›h
›tdz’
ðztopzsfc
�i2T
Mih dz1 r
0
›h
›tZ ,
(7)
FIG. 7. (left) Logarithm of the joint distribution of vertical velocity and MSE anomaly at 531m. The black
contour corresponds to the isopleth 21 of the logarithmic distribution. (right) Sinks of MSE at the top of the
subcloud layer divided by three transport categories: updraft, downdraft, and environment. The blue (green) bars
show the rates according to the broad (narrow) definition of the categories introduced in Thayer-Calder and
Randall (2015), whereas the red bars show the results obtained using the definition we have given using Lagrangian
particles.
846 JOURNAL OF THE ATMOSPHER IC SC IENCES VOLUME 73
where Z5 ztop 2 zsfc and h is the average MSE per unit
area in the subcloud layer. This can be further approx-
imated by substituting h in the integral with its value at
the top of the subcloud layer htop to give
›[rh]
›t’Z
�i2T
Mihtop
1 r0
›h
›t
!. (8)
Substituting Eq. (8) into Eq. (4), we can write a budget
equation for h:
›h
›t5
1
Z�i2T
wi(h
i2 h
top)1
SH1LH
r0Z
1QR, (9)
where wi is the velocity associated with the mass fluxMi
and we have substituted the term for the radiative ef-
fects with its subcloud-layer average QR. We have de-
rived this formula for an atmospheric column of unit
area, but a generalization to an extended domain such as
the one we are using is straightforward.
The choice of substituting h with htop in the right side
of Eq. (7) may have seemed a bit arbitrary, but it can be
fully appreciated in light of Eq. (9), where it is used as a
reference to determine the anomalies of MSE at the top
of the subcloud layer: both terms hi and htop are com-
puted at the same height.
Another source of arbitrariness that we have deliber-
ately left vague in our derivation is the definition of the
three transport categories. As is well known, the lines
between the three categories are blurry enough to allow
for multiple ways to address this issue. For example,
Thayer-Calder and Randall (2015) used two different
definitions to compute the budget ofMSEat the top of the
boundary layer. In the first one, called broad, each grid
box at 500-m height is considered a downdraft (updraft)
box if it contains any cloud ice, water, or precipitation and
has a negative (positive) velocity; if the box does not
contain any condensate, it is considered as environment.
In the second, called narrow, a grid box is considered a
downdraft (updraft) box if the vertical velocity is
below21ms21 (above 1ms21) for two continuous levels,
above and below 500m, and if it contains any condensate;
all other grid boxes are considered environment.
A potential problemwith this approach is that some of
the properties of a parcel, like its vertical velocity or its
liquid water specific humidity, near the top of the
boundary layer or cloud base are not necessarily in-
dicative of what transport category the parcel belongs
to. For instance, a parcel that will enter a convective
updraft might have an LCL at a higher altitude than
500m, or it might encounter significant convective in-
hibition, which will then sensibly reduce its vertical ve-
locity, before reaching its LFC. A similar story could
also hold true for parcels in downdrafts. We have tried
reproducing the results of Thayer-Calder and Randall
(2015) at various heights around 500m and found the
conclusions, particularly the contribution by updrafts, to
change according to which height is being used.
To remove this ambiguity, we propose a slightly dif-
ferent approach using the Lagrangian particles. Let us
go back to the joint distribution of vertical velocity and
MSE anomaly shown in Fig. 7 (left). Whenever a par-
ticle reaches the top of the subcloud layer, its vertical
velocity and the future 60min of its history are consid-
ered. If the particle has positive vertical velocity and,
within the specified time frame, acquires positive
buoyancy at an altitude higher than 1.5 km, the particle
is considered an updraft particle; if the particle has
negative vertical velocity and enters a grid box with a
density potential temperature anomaly lower than20.5K
in less than an hour, the particle is considered a downdraft
particle. If none of the above conditions are true, the
particle is considered an environment particle.
In Fig. 7 (right), we show the contributions to the budget
of MSE by the three transport categories. To compare our
method with those of Thayer-Calder and Randall (2015),
we measure the fluxes of MSE at 531m, which, in our
model, is the height closest to 500m and also corresponds
to the top of the subcloud layer. The blue (green) bars
refer to the results obtained using the broad (narrow)
definitions explained above, whereas the red bars refer to
the method using Lagrangian particles explained above.
Like Thayer-Calder and Randall (2015), we find turbulent
mixing in the environment to be the biggest contributor to
the total MSE sinks, although the contribution from up-
drafts and downdrafts is not negligible. The export of high
MSE air by convective updrafts provides a sink of MSE of
2.43Kday21, which is 16% of the total, amounting to
15.51Kday21, whereas the low-MSE air injected in the
subcloud layer by precipitation-driven downdrafts reduces
MSE at the rate of 1.81Kday21, 12% of the total. As
documented in the supplemental information, we have
also tested the sensitivity of the results to the sampling
height and found that our conclusions are robust.Of all the
heights where we have tested our results, 531m is the one
where the three methods are closest to each other, espe-
cially in predicting the contribution by downdrafts. Finally,
we want to note that the surface sensible and latent heat
fluxes contribute to the budget ofMSEwith a source equal
to 16.28Kday21, whereas radiative effects provide a sink
of 0.86Kday21.
c. On the maintenance of precipitation-drivendowndrafts
The next question that we want to focus on in the
present work is how to quantify the importance of rain
FEBRUARY 2016 TORR I AND KUANG 847
evaporation and whether it is bigger than that of con-
densate loading in maintaining precipitation-driven
downdrafts. Because we are concerned with the role of
rain evaporation from the dynamical point of view, we
propose using a purely dynamical quantity such as work
to address the problem.
The buoyancy of every particle can be defined in
terms of its density potential temperature:
b5 g
ur2 u
r
ur
!, (10)
where g is the gravitational constant and ur is the hori-
zontal average of density potential temperature.
At each time step, the contribution to a particle’s
buoyancy by the condensate loading bl is given by
bl52g
qlu
ur
!, (11)
from which it follows that the work done by condensate
loading on a particle in a precipitation-driven downdraft
can be defined as
Wl52g
ðzizcore
qlu
ur
!dz , (12)
where the integral is taken from the initial height zi to
the height where the particle enters the core of a cold
pool zcore.
To do the same for rain evaporation, we consider the
change in the thermodynamic state of a particle at each
time step due to the evaporation of a certain amount of
precipitation qev. Assuming that the evaporation takes
place at the end of each time step and that the MSE of a
particle is left invariant, the particle’s potential tem-
perature will change by
uev52
Ly
cp
qev, (13)
where Ly is the latent heat of vaporization, equal to
2.5104MJkg21 at 08C, and cp is the specific heat of dry
air at constant pressure, equal to 1004 J kg21K21. This
change in thermodynamic properties implies a change in
the particle’s buoyancy that, after a bit of algebra, can be
written as follows:
bev52
g
ur
"L
y
cp
(11 «qy2q
l)2 «u
#qev. (14)
Much in the same spirit of Eq. (12), we define the work
done on a particle by rain evaporation as
Wev52g
ðzizcore
"L
y
cp
(11 «qy2 q
l)2 «u
#qev
ur
dz . (15)
We will follow the convention that work done on a
particle by the environment is positive.
The definition we have given above will provide us
with a way to quantify the importance of each mecha-
nism in maintaining a downdraft. Considering again the
Lagrangian particles we sampled earlier to construct
Fig. 2, we compute the two types of work for each par-
ticle and average together the values for particles with
the same initial height with the assumption that they
have a comparable history and experienced the same
phenomena. Figure 8 shows a comparison of the average
work done on particles with different initial heights by
condensate loading (solid red) and by rain evaporation
(solid blue). Notice that, apart from the bottom 300m,
the contribution due to evaporation is smaller than that
due to loading by a factor of approximately 0.4.
We can test the consistency of this result by using an
Eulerian approach. First, we invoke some approxima-
tion to simplify Eqs. (12) and (15). To first order, we can
exchange ur with u in the denominator, and we can
consider qy and ql to be much less than 1. Then, since «u
is more than an order of magnitude smaller than the ratio
Ly/cp, we can neglect the second term in square brackets
in Eq. (15). With these simplifications, we can write
FIG. 8. Comparison between the total work done on Lagrangian
particles with different initial heights by different contributions to
buoyancy (solid lines) and total buoyancy (dashed lines). Line
colors refer to the contributions by condensate loading (red) and
the contribution by rain evaporation (blue).
848 JOURNAL OF THE ATMOSPHER IC SC IENCES VOLUME 73
Wev
Wl
’
L
y0
cpQ
!ð qevdzð
qldz
, (16)
where Ly has been approximated by its value at 08C,2.5104MJkg21, and Q represents a scale for the poten-
tial temperature and equals 300K. To evaluate the in-
tegrals in the above equation, we will consider a parcel
descending from an altitude of 1 km, the location of the
peak in Fig. 2. For every height, we compute the distri-
bution of negative vertical velocities of grid boxes where
qp is nonzero, and we assume that the vertical velocity of
the parcel we are considering equals the median of the
distribution. This construction is to ensure that the
parcel is representative of parcels within precipitation-
driven downdrafts.
The integral in the denominator is proportional to the
average value of liquid water content of the parcel, and
it can be quickly put in discrete form as
ðzizsfc
qldz5 �
ki
i5ksfc
(ql)iDz
i, (17)
whereDzi is the vertical grid spacing at height zi, and ksfcand ki are the model levels corresponding to heights zsfcand zi, respectively.
The integral in the numerator requires a little more
care. The integrand represents the specific humidity of
liquid water that is evaporated into the parcel as it
travels a distance equal to dz. By definition, this can be
rewritten as
qev5 (dqsrc
y ) dt5 (dqsrcy )
dz
w, (18)
where (dqsrcy ) is the rate of generation of water vapor
by rain evaporation and dt is the time it takes the
parcel to go from z1 dz to z. It follows that we can
evaluate the integral in the numerator of Eq. (16) by
computing the sum:
ðzizsfc
qevdz5 �
ki
i5ksfc
(dqsrcy )
i
�Dz2iw
i
�. (19)
The liquid water specific humidity and the evaporation
rate can be diagnosed from the model outputs, and their
values in the bottom 5km can be seen in Fig. 5. Plugging
in all the numbers, we can compute that the ratio be-
tween the work done by rain evaporation and that done
by condensate loading is 0.36, which compares reason-
ably well with the estimate of 0.41 that is obtained using
the Lagrangian particles.
As discussed by Davies-Jones (2003), buoyancy in a
nonhomogeneous environment, such as the one we are
examining, is an ill-defined concept. Instead, what
should really be considered as the effective buoyancy
experienced by a parcel is the sum of the Archimedean
buoyancy, which we have defined in Eq. (1), and the
buoyancy-related pressure gradient, which cannot be
determined locally. Recently, it has been shown (Torri
et al. 2015; Jeevanjee and Romps 2015) that parcels that
have been triggered by cold pools and that are ascending
toward their LFC are subject to a significant cancellation
between buoyancy and buoyancy-related pressure gra-
dients. This leads us to ask whether our conclusions on
the minor role played by rain evaporation in maintain-
ing the downdraft are robust when buoyancy-related
pressure gradients are taken into account. To address
this point, for simplicity, we first assume that the con-
tributions to the buoyancy-related pressure gradients at
every moment can be viewed as approximately pro-
portional to the contributions to the Archimedean
buoyancy by a factor given by the ratio of the buoyancy-
related pressure gradient and buoyancy, which we will
refer to as buoyancy ratio b:
b 5def ›
zpb
b’›zpb,l
bl
’›zpb,ev
bev
, (20)
where pb is the buoyancy-related pressure perturbation
and pb,l and pb,ev are its contributions due to condensate
loading and rain evaporation, respectively.
In this way, we can reconstruct the various contribu-
tions to the buoyancy-related pressure gradients very
simply, and the calculation of the work done by these
contributions is straightforward.
The two contributions to the work done by total
buoyancy—bywhichwemean the sumof theArchimedean
buoyancy and the buoyancy-related pressure gradients
(Torri et al. 2015)—are shown by the dashed lines in
Fig. 8, and they suggest that our conclusions are robust,
even when buoyancy-related pressure gradients are
taken into account. A close inspection of the vertical
profiles of buoyancy ratio (not shown) suggests that this
quantity tends to be maximum in magnitude in the
subcloud layer, with values approaching 21, and be-
comes closer to 0 with increasing height. This explains
why the particles that seem most affected by the cor-
rection are those with an initial height smaller than 1km.
A natural question that arises now is whether con-
densate loading dominates over rain evaporation at any
moment during the particle’s descent in the downdraft
or if the two mechanisms are important at different
stages. To investigate this, instead of considering an in-
tegrated quantity such as the work, we look directly at
FEBRUARY 2016 TORR I AND KUANG 849
the contributions to buoyancy given by the two mecha-
nisms. We present the results in Fig. 9, with Fig. 9 (left)
referring to the contribution from condensate loading
and Fig. 9 (right) referring to that from rain evaporation.
As for the construction of Fig. 8, we use the initial
height of a particle as a sampling criterion: the histories
of particles with the same initial height are averaged
together, and the results are presented for each value of
initial height. Thus, each column in Fig. 9 represents the
vertical profile of the average contribution to the
buoyancy of particles having an initial height given by
the abscissa of the column in the figure.
In comparing the two plots, one should be careful to
notice that the color bars differ by a factor of 2.5. This
was done to make it possible for the eye to easily dis-
tinguish the main features of each contribution at every
height for different types of particles. Therefore, in spite
of the immediate appearance, condensate loading con-
tributes to buoyancy more than evaporation almost ev-
erywhere, except for very near the surface, where both
contributions seem of equal magnitude.
Another interesting feature of the two plots is that, for
any initial height, the altitude where each contribution is
maximum is different: for loading, the peak tends to be
in the initial stages of the downdraft, whereas for
evaporation the maximum is in the subcloud layer.
For the condensate loading, the peak is essentially due
to two factors: the first is that rain columns are not steady
but, rather, quickly grow to a maximum and then decay
over time; the second is the fact that precipitating water
in a rain column flows through the Lagrangian particles
or, in other words, the vertical velocity of the particles is
typically smaller than the terminal velocity of raindrops.
To check that the latter is true, we can consider
Fig. 10, which shows the average vertical velocity
experienced by particles of various initial heights at
different stages of their descent. Above the bottom few
hundred meters, where the hydrostatic pressure per-
turbation in the core of cold pools causes the downdraft
to slow down, the vertical velocities experienced by
the descending particles are roughly between 20.5
FIG. 9. Vertical profiles of the time-averaged change of buoyancy due to (left) condensate loading and (right) rain
evaporation for Lagrangian particles in precipitation-driven downdrafts with different initial heights.
FIG. 10. Vertical profiles of time-averaged vertical velocities for
Lagrangian particles in precipitation-driven downdrafts with dif-
ferent initial heights.
850 JOURNAL OF THE ATMOSPHER IC SC IENCES VOLUME 73
and 21.5m s21. Considering the precipitation rate and
the precipitating liquid water content in the grid boxes
of the bottom 3km of the model, we can estimate that
the average terminal velocities of raindrops are between
25 and 26ms21, well outside the range of average ve-
locities of Lagrangian particles within precipitation-driven
downdrafts. As a side note, we want to point out that the
terminal velocities diagnosed from the model are consis-
tent with data from observations: the terminal velocities
for raindrops with diameters between 1 and 5mm go
from 24.03 to 29.09m s 21 (Gunn and Kinzer 1949).
To understand why the contribution to buoyancy by
rain evaporation—and, more generally, the evaporation
rate—peaks at low altitudes, we must take a step back
and think about how liquid water evaporates in a par-
ticle in the downdraft. It is well known (see, e.g.,
Kamburova and Ludlam 1966; Betts and Silva Dias
1979) that evaporation of a falling raindrop of radius
r and mass m can be modeled as a diffusion process:
Dm
Dt5 4pDC
yrr(q
y2 q
sat) , (21)
whereD is the diffusion coefficient of water vapor in air,
Cy is a ventilation factor, r is the density of the parcel,
and qsat is the saturation specific humidity. If we consider
an air parcel with volume V and assume, for simplicity,
that it contains N raindrops of equal radius R, then the
liquid water content of the parcel can be simply related
to the radius of the raindrop by the following:
ql5
�4pr
lN
3rV
�R3 5
def�4pDC
y
VK
�3
R3 , (22)
where rl is the density of liquid water, and, for simplicity,
we have defined a new variable K as
K5DCy
�48p2r
NV2rl
�1/3
. (23)
With some simple algebra, it is easy to show that Eq. (21)
can be rewritten as
Dql
Dt5Kq1/3
l (qy2 q
sat) . (24)
Figure 11 shows the average product of the cubic root of
the cloud liquid water and the saturation deficit for
particles with different initial heights. Assuming that K
is constant at all heights and for particles from all initial
heights, the figure can be compared directly with the
contribution to buoyancy by rain evaporation. Notice
that, for each initial height, the profiles show a peak in
the subcloud layer, roughly at the same height where the
contribution by evaporation is maximum. Thus, one
could say that the peak in evaporation rate in the sub-
cloud layer is the result of two factors: the saturation
deficit, which increases as the surface is approached (not
shown), and the liquid water content of each parcel,
which, as can be inferred from Fig. 9 (left), decreases in
the latter stages of the downdraft.
Given the crude assumptions that we have made to
derive Eq. (24), it is not surprising that the agreement
between Fig. 11 and Fig. 9 (right) is only partial: unlike
the profiles of the contribution by evaporation, the
magnitude of the product seems generally to decrease
for particles with increasing initial height. We expect
that accounting for a distribution of radii and by allow-
ing the number of raindrops N to vary with height and
initial height would improve the agreement. However,
we want to stress that this mismatch does not change our
conclusions.
4. Conclusions
We have used an LPDM to address two important
aspects regarding the dynamics of precipitation-driven
downdrafts: namely, their initial height and the role
played by rain evaporation in maintaining them. By
sampling all the Lagrangian particles that are injected in
the coldest regions of cold pools in the subcloud layer by
downdrafts, we have determined that the vast majority
of such particles start descending from a height lower
FIG. 11. Profiles of the time average of the cubic root of liquid
water specific humidity multiplied by the saturation deficit for
Lagrangian particles in precipitation-driven downdrafts.
FEBRUARY 2016 TORR I AND KUANG 851
than 2.5 km in the model domain. Looking at the posi-
tion of the particles at times prior to their entrance in a
precipitation-driven downdraft, we have shown that
most of them originate in the subcloud layer; almost
none come from high altitudes.
As a further check, we have broadened our sampling
criterion and considered all particles that are part of
downdrafts at any height, regardless of whether they will
be injected in a cold pool or not. With this, we have
shown that downdrafts exist at all heights in the
model but a few hundred meters past their initial
height. This provided additional support to the claim
that precipitation-driven downdrafts originate at very
low altitudes and do not, therefore, bring very cold and
dry air into the subcloud layer.
This led us to the question of what balances the sur-
face fluxes and keeps the boundary layerMSE relatively
steady in time. Using a definition of the three transport
categories—updraft, downdraft, and environment—
based on the Lagrangian particles, we have concluded
that most of theMSE sink is to be attributed to turbulent
mixing or wave activity in the environment, although the
contributions by updrafts and downdrafts are non-
negligible, their sum accounting for 28% of the total.
We then proceeded to quantify the role of rain
evaporation in maintaining the precipitation-driven
downdrafts. By looking at the total work done on each
Lagrangian particle that is injected in the core of a cold
pool by a downdraft and distinguishing the contribution
due to condensate loading from that due to rain evap-
oration, we showed that the latter is smaller than the
former, roughly by a factor of 0.4.
Considering the change of the Lagrangian particles’
buoyancy due to condensate loading and rain evapora-
tion, we showed that the former is bigger than the latter
at all heights and tends to bemaximum in the first part of
the downdraft descent; on the other hand, the latter is
highest in the second part of the descent, when the
particles are in the subcloud layer.
We interpreted the first effect as due to the fact that
the terminal velocity of rain droplets is higher in mag-
nitude than the vertical velocity of the descending
Lagrangian particles and that the liquid water content as-
sociated with each rain shaft is not constant in time.
As for rain evaporation, we have argued that its
maximum in the subcloud layer is simply the combination
of saturation deficit and the liquid water specific humidity,
which enters the equation for evaporation rates only in a
cubic root. In particular, even though the precipitation
experienced byLagrangian particles is largest in the initial
stages of the downdraft, the small saturation deficit does
not allow for large amounts of rain to be evaporated.Only
in the subcloud layer is the saturation deficit large enough
to make the contribution of rain evaporation in main-
taining the downdraft more significant.
Since the dynamics of downdraft maintenance is
tightly related to microphysical processes such as rain
evaporation, it seems natural to ask how sensitive our
conclusions are to the microphysics scheme. For this
reason, we have run the same simulation described in
section 2 using the two-moment microphysics scheme
introduced inMorrison et al. (2005) and have carried out
the same analysis described in the previous section.
Apart from some slight differences, we found that the
conclusions of this study are robust.
Although this robustness gives us some confidence
over the generality of our findings, we want to stress that
we have only considered an oceanic case in RCEwith no
wind shear and, a priori, we do not expect our results to
hold for other cases. For instance, it is very well possible
that, in a continental case with a much drier boundary
layer, evaporation of rain will play a much bigger role in
maintaining the downdraft. This could also be the case in
the presence of wind: a considerable vertical wind shear
might change the geometry of a precipitating cloud,
exposing a larger part of a precipitating column to an
unsaturated environment.
Finally, the conceptual framework that emerges
from this manuscript can have important consequences
for climate models. In particular, our findings suggest
that the premises on which the BLQE hypothesis is
founded—that downdrafts are responsible for most of
the MSE sink in the boundary layer—may not be real-
ized in nature. Although some models have been shown
to better reproduce certain features of the tropical cli-
mate, such as the MJO, when strong convective down-
drafts are included (Mishra and Sahany 2011), we suggest
that the key for amore accurate simulation of the tropical
atmosphere is in improving parameterizations of the
transport due to turbulent mixing or wave activity in the
environment. Mesoscale organization of convective sys-
tems, absent in this study, may also affect the results.
Notwithstanding these additional scenarios, which clearly
warrant further investigations, our analysis of the oceanic
RCE case addresses a basic scenario that is potentially
representative of large areas over the tropical oceans with
modest wind shears.
Although they play only a secondary role in the MSE
budget of the tropical boundary layer, downdrafts still
remain an important component of convective systems
that should be included in convective parameterizations.
Our results could also be useful for this task: first, they
indicate the initial height tends to be in the lower tro-
posphere; second, the fact that rain evaporation contrib-
utesmuch less than condensate loading inmaintaining the
downdraft indicates that downdrafts might be more
852 JOURNAL OF THE ATMOSPHER IC SC IENCES VOLUME 73
sensitive to the model precipitation rate rather than to the
relative humidity.
Acknowledgments. The authors thank Marat
Khairoutdinov for access to the SAM code and Paul
Edmon for precious and tireless assistance with the
HarvardOdyssey cluster on which the simulations were
run. This research was partially supported by the Office
of Biological and Environmental Research of the U.S.
Department of Energy under Grant DE-SC0008679 as
part of the ASR program and by NSF Grants AGS-
1062016 and AGS-1260380.
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